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Ornithosuchidae is an extinct family of
pseudosuchian Pseudosuchia is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely related to crocodilians than to birds. Pseudosuchians are also informally known as "crocodilian-line archosaurs". Prior to ...
archosaur Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avian ...
s (distant relatives of modern
crocodilians Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period (Cenomanian stage) and are the closest living r ...
) from the
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
period. Ornithosuchids were
quadrupedal Quadrupedalism is a form of locomotion where four limbs are used to bear weight and move around. An animal or machine that usually maintains a four-legged posture and moves using all four limbs is said to be a quadruped (from Latin ''quattuor ...
and facultatively
bipedal Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' 'double' ...
(e.g. like chimpanzees), meaning that they had the ability to walk on two legs for short periods of time. They had distinctive, downturned snouts, unique, "crocodile-reversed" ankle bones, and several other features that distinguish them from other archosaurs. Ornithosuchids were geographically widespread during the
Carnian The Carnian (less commonly, Karnian) is the lowermost stage of the Upper Triassic Series (or earliest age of the Late Triassic Epoch). It lasted from 237 to 227 million years ago (Ma). The Carnian is preceded by the Ladinian and is followed by ...
and
Norian The Norian is a division of the Triassic Period. It has the rank of an age (geochronology) or stage (chronostratigraphy). It lasted from ~227 to million years ago. It was preceded by the Carnian and succeeded by the Rhaetian. Stratigraphic ...
stages of the Late Triassic with members known from
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
, Brazil, and the United Kingdom. Four genera, comprising ''
Ornithosuchus ''Ornithosuchus'' (meaning "bird crocodile") is an extinct genus of pseudosuchians from the Late Triassic ( Carnian) Lossiemouth Sandstone of Scotland. It was originally thought to be the ancestor to the carnosaurian dinosaurs (such as ''Allosau ...
'', '' Venaticosuchus, Dynamosuchus,'' and '' Riojasuchus'' are presently known. The family was first erected by German paleontologist Friedrich von Huene in 1908.


Description


Skull

Ornithosuchids can be identified by the presence of an arched
diastema A diastema (plural diastemata, from Greek διάστημα, space) is a space or gap between two teeth. Many species of mammals have diastemata as a normal feature, most commonly between the incisors and molars. More colloquially, the condition ...
, a gap between the teeth at the front of the snout. When the jaw is closed, two large curved
dentary In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
(lower jaw) teeth fit into the diastema, which is positioned between the
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has ...
and maxilla. There are two shallow depressions on the wall of the diastema to accommodate these teeth. The large dentary teeth of ''Ornithosuchus'' and ''Riojasuchus'' are placed behind a smaller procumbent dentary tooth that sticks out from the jaw. This type of tooth position is not seen in any other basal archosaurs. Another characteristic feature of ornithosuchids is their unusual downturned, overhanging snout, seen in ''Riojasuchus'' and ''Venaticosuchus'', but not ''Ornithosuchus''. Several other features distinguish ornithosuchids from all other early archosaurs. ''Ornithosuchus'' and ''Riojasuchus'' both possess a small fenestra, or hole, between the palatine and
pterygoid bone The pterygoid is a paired bone forming part of the palate of many vertebrates Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and ...
s of the
palate The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly sepa ...
, i.e. the roof of the mouth. The contact between the nasal and
prefrontal bone The prefrontal bone is a bone separating the lacrimal and frontal bones in many tetrapod skulls. It first evolved in the sarcopterygian clade Rhipidistia, which includes lungfish and the Tetrapodomorpha. The prefrontal is found in most mo ...
s of the skull is small or absent, excluded by a large contact between the frontals and
lacrimals The lacrimal bone is a small and fragile bone of the facial skeleton; it is roughly the size of the little fingernail. It is situated at the front part of the medial wall of the orbit. It has two surfaces and four borders. Several bony landmarks of ...
. In other archosaurs, including
rauisuchian "Rauisuchia" is a paraphyletic group of mostly large and carnivorous Triassic archosaurs. Rauisuchians are a category of archosaurs within a larger group called Pseudosuchia, which encompasses all archosaurs more closely related to crocodilians ...
s,
aetosaur Aetosaurs () are heavily armored reptiles belonging to the extinct order Aetosauria (; from Greek, (aetos, "eagle") and (, "lizard")). They were medium- to large-sized omnivorous or herbivorous pseudosuchians, part of the branch of archosaurs ...
s,
pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 to ...
s, and
dinosauromorph Dinosauromorpha is a clade of avemetatarsalian archosaurs (reptiles closer to birds than to crocodilians) that includes the Dinosauria (dinosaurs) and some of their close relatives. It was originally defined to include dinosauriforms and lage ...
s, the nasal-prefrontal contact separates the frontal from the lacrimal.


Postcranial skeleton

The postcranial skeleton is nearly completely known in ''Riojasuchus'', incomplete in ''Ornithosuchus'', and entirely unknown in ''Venaticosuchus''. As a result, it is uncertain whether all of the postcranial traits seemingly unique to ornithosuchids actually occurred in all members of the family. Ornithosuchids known from decent postcranial remains typically had about 9
cervical In anatomy, cervical is an adjective that has two meanings: # of or pertaining to any neck. # of or pertaining to the female cervix: i.e., the ''neck'' of the uterus. *Commonly used medical phrases involving the neck are **cervical collar **cervic ...
(neck), 14-15 dorsal (back), 3
sacral Sacral may refer to: *Sacred, associated with divinity and considered worthy of spiritual respect or devotion *Of the sacrum The sacrum (plural: ''sacra'' or ''sacrums''), in human anatomy, is a large, triangular bone at the base of the spin ...
(hip), and over 20 caudal (tail)
vertebrae The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic i ...
. Above each vertebra was a pair of bony scutes known as osteoderms. The femur (thigh bone) has a pronounced anterior trochanter. The anterior trochanter, sometimes known as the "lesser trochanter" (but unrelated to the lesser trochanter of the femur in humans), is a ridge on the outer surface of the femur, near the
femoral head The femoral head (femur head or head of the femur) is the highest part of the thigh bone ( femur). It is supported by the femoral neck. Structure The head is globular and forms rather more than a hemisphere, is directed upward, medialward, and a ...
. It was probably an insertion point for the ''iliofemoralis cranialis'' muscle which helps to raise the leg. Most archosaurs and archosaur relatives lack a distinct anterior trochanter, but ornithosuchids are an exception, along with most dinosauromorphs (dinosaurs and their close relatives). Much like the femur, the fibula (outer shin bone) also has a distinctive point for muscle insertion. The muscle in question is the ''iliofibularis'', which helps to straighten the limbs. In most archosaurs, the ''iliofibularis'' is inserted onto the fibula by means of a tiny ridge on the proximal part of the fibula, near the knee. However, ornithosuchids have a much larger, knob-shaped ''iliofibularis'' insertion point located about midway down the shaft of the fibula. Phytosaurs and aetosaurs also share a knob-like attachment point midway down the fibula, so it is unclear whether the case in ornithosuchids is a unique case of convergent evolution, or alternatively the retention of a trait independently lost by several archosaur lineages. Unlike most other early archosaurs, the pedal
ungual An ungual (from Latin ''unguis'', i.e. ''nail'') is a highly modified distal toe bone which ends in a hoof, claw, or nail. Elephants and ungulates have ungual phalanges, as did the sauropod Sauropoda (), whose members are known as sauropods (; ...
s (the distalmost bones of the feet that form claws) are laterally compressed. They are sharp and recurved. The unguals are very deep, being taller than they are long, especially on the inner digits. This type of claw is not seen in any other Triassic archosaur except for pterosaurs. Major archosaur groups have often been distinguished from each other based on the structure of their ankles. In most crurotarsans, the
astragalus ''Astragalus'' is a large genus of over 3,000 species of herbs and small shrubs, belonging to the legume family Fabaceae and the subfamily Faboideae. It is the largest genus of plants in terms of described species. The genus is native to tempe ...
has a convex projection that fits into a concave space in the
calcaneum In humans and many other primates, the calcaneus (; from the Latin ''calcaneus'' or ''calcaneum'', meaning heel) or heel bone is a bone of the tarsus of the foot which constitutes the heel. In some other animals, it is the point of the hock. S ...
. This condition is often referred to as a "crocodile-normal" ankle, as it is the most common ankle type in crurotarsans. Ornithosuchids are unique among crurotarsans, and all other archosaurs, in their possession of a "crocodile-reversed" ankle. In a "crocodile-reversed" ankle, the placement of the concavity is reversed: instead of being on the calcaneum, it is on the astragalus. In ornithosuchids, the calcaneum bears a convex projection that is analogous to the convex projection on the "crocodile-normal" astragalus.


Phylogeny

Ornithosuchidae is generally considered to be within the larger clade Suchia, which includes aetosaurs, rauisuchians, and crocodylomorphs. Below is a cladogram based on Nesbitt (2011), showing the placement of Ornithosuchidae in Archosauriformes.


References


External links


Palaeos Vertebrates
{{Taxonbar, from=Q134276 Late Triassic pseudosuchians Late Triassic first appearances Rhaetian extinctions Prehistoric reptile families